LARVAL ECOLOGY OF MARINE BENTHIC INVERTEBRATES: PALEOBIOLOGICAL IMPLICATIONS

Abstract

Summary1. Modes of larval development play important roles in the ecology, biogeography, and evolution of marine benthic organisms. Studies of the larval ecology of fossil organisms can contribute greatly to our understanding of such roles by allowing us to race effects on evolutionary time scales.2. Modes of development can be inferred for well preserved molluscan fossils because the size of the initial larval shell (Protoconch I in gastropods, Prodissoconch I in bivalves) reflects egg size. Other morphological criteria are also available, and a comparative approach based on related taxa with known development may be the most reliable method. By combining larval and adult traits, it is possible to recognize modes of larval development in at least some fossil bryozoans, brachiopods, and echinoderms as well. (a) Planktotrophic larvae arise from small eggs, are released in enormous numbers with little parental investment per offspring, and suffer tremendous mortality during and shortly after a planktic existence. These larvae feed on the plankton during development, and are commonly capable of a prolonged free‐swimming existence, and thus wide dispersal. (b) Nonplanktotrophic larvae (which include both planktic lecithotrophic forms and ‘direct developers’) generally arise from large eggs, with relatively few young produced per parent. Relative to planktotrophic larvae, nonplanktotrophic larvae generally receive greater parental investment per larva, and larval mortality is generally lower. These larvae rely on yolk for nutrition during development, and planktic durations are generally much briefer than for species with planktotrophic larvae, so that dispersal capability is considerably less. Energetic investment per egg is generally higher than in planktotrophs, but as there are lower fecundities as well it is difficult to generalize about the total energetic cost of one mode of reproduction against the other.3. Owing to the high dispersal capability of planktotrophic larvae, it has been suggested that species with such larvae will be geographically widespread, geologically long‐ranging, and exhibit low speciation and extinction rates. Species with nonplanktotrophic larvae will tend to be geographically more restricted, geologically short‐ranging, and exhibit high speciation and extinction rates (again, as a consequence of their characteristically low larval dispersal capabilities).4. Recognition of differential dispersal capabilities can play a role in paleobiogeo‐graphic analyses. Concurrent study of the distribution of groups with contrasting modes of development will permit testing of hypotheses concerning timing, magnitudes and frequencies of migration and vicariance events.5. Larval types are not randomly distributed in the oceans, but relationships with other aspects of the organisms' biology and habitats are very complex. Mode of development varies with: (a) Ecology. A simple r–––K model of adaptive strategies is clearly insufficient to explain the observed relationships: while many ‘equilibrium’ species have nonplanktotrophic larvae, and organisms living in less prdictable environments often have planktotrophic larvae, some of the most opportunistic marine species have nonplanktotrophic larvae. Nonetheless, planktotrophic development seems most suited for exploitation of patchy but widespread habitats. (b) Latitude. At shelf depths, planktotrophy is predominant in the tropics, and decreases sharply at high latitudes. (c) Depth. Incidence of planktotrophy decreases with depth across the continental shelf, at least in some taxa. Beyond the shelf, many deep‐sea organisms are nonplanktotrophic (e.g. most bivalves, peracarid crustaceans), but planktotrophic development appears to be present in other groups (prosobranch gastropods, ophiuroids, and bivalves inhabiting transient habitats such as sunken wood and hydrothermal vents).These trends in developmental types will be accompanied by trends in evolutionary rates and patterns as outlined above. The study of larval ecology by paleobiologists will yield insights into the processes that gave rise to ancient evolutionary and biogeographic patterns, and will permit the development and testing of hypotheses on the origins of the patterns observed in modern seas.